Zoosyst. Evol. 99 (2) 2023, 489-501 | DOI 10.3897/zse.99.102345 eee BERLIN Identification of the rare deep-dwelling goby Suruga fundicola Jordan & Snyder, 1901 (Gobiiformes, Gobiidae) from the Yellow Sea Changting An’, Ang Li!:*, Huan Wang?*, Busu Li!*, Kaiying Liu!, Hongyue Sun!, Shufang Liu’, Zhimeng Zhuang?, Richard van der Laan? 1 National Key Laboratory of Mariculture Biobreeding and Sustainable Goods, Yellow Sea Fisheries Research Institute, Chinese Academy of Fishery Sciences, 106 Nanjing Road, Qingdao, Shandong, China 2 Laboratory for Marine Fisheries Science and Food Production Processes, Laoshan Laboratory, Qingdao, China 3 Grasmeent 80, 1357JJ Almere, Netherlands https://zoobank. org/BED03B87-4DB0-4940-BC16-A 1S86CC92EF7 Corresponding author: Shufang Liu (liusf@ysfri.ac.cn) Academic editor: Nalani Schnell # Received 19 February 2023 # Accepted 13 September 2023 @ Published 5 October 2023 Abstract During the 2022 R/V cruises in the Yellow Sea, four goby specimens (51.2—-63.5 mm) were captured by using an Agassiz trawl at a water depth of about 70 meters. These specimens were identified as Suruga fundicola, mainly by the morphometric characters. Their identification was further confirmed by a molecular phylogenetic analysis based on /2S and CO/ mtDNA genes. Considering that the four specimens were in good condition and that the original description is brief, a detailed description of the specimens is given. Moreover, the present study presents a preliminary analysis of its phylogenetic position within the Acanthogobius-lineage (Gobiidae). The discovery of this goby in the Yellow Sea enriches our knowledge of the fish diversity and distribution of this region, and sheds some light on the ecological habitat of these gobies. Key Words Acanthogobius-\ineage, distribution, morphology, mtDNA genes, species identification, taxonomy Introduction The gobies (order Gobiiformes) include about 2400 spe- cies divided into about 320 genera, which are widely distributed throughout the tropical, subtropical, and tem- perate regions of the world (Parenti 2021; Fricke et al. 2023). These goby species are known from the deep see waters (at a depth of over 300 meters) to elevated moun- tain streams (at an altitude of over 1000 meters) (Iwata et al. 2000; Shibukawa and Aonuma 2007; Parenti 2021). About 90% of them dwell in marine water, and generally live at the bottom of the sea with a depth of no more than 30 m (Wu and Zhong 2008; Akihito et al. 2013). Only a few species of goby inhabit deep waters; examples from the Western Pacific are species of the genera Obliquogo- bius Koumans, 1941 and Suruga Jordan & Snyder, 1901 (Shibukawa and Aonuma 2007; Fujiwara and Shibukawa 2022). According to the record, Suruga fundicola Jordan & Snyder, 1901 can inhabit the bottom of deep water off the coast (depth about 40-400 meters; Akihito et al 2013; Choi and Lee 2019). The goby genus Suruga Jordan & Snyder, 1901 of the family Gobiidae comprises only one species, Suruga fundi- cola Jordan & Snyder, 1901 (Akihito et al 2013; Fricke et al. 2023). Based on axial skeletal features, Swruga was placed in the Acanthogobius-group (sometimes denoted as the tribe Acanthogobiini Parenti 2021) with 7 other genera, Acanthogobius Gill, 1859, Amblychaeturichthys Bleeker, 1853, Chaeturichthys Richardson 1844, Lophiogobius Gunther, 1873, Pterogobius Gill, 1863, Sagamia Jordan & Snyder, 1901 and Siphonogobius Shibukawa & Iwata, 1998 (Birdsong et al. 1988; Shibukawa 1997). Copyright Changting An et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 490 Changting An et al.: Identification of the Suruga fundicola from the Yellow Sea These eight genera including 18 species, are defined by a unique pattern of the dorsal-pterygiophore formula 3/ I If 1 II I O (indicating the relationship between the pterygiophores of the dorsal fins and the corresponding spines of the vertebrae), and these genera are regarded as a putative monophyletic assemblage (Akihito et al. 1984; Shibukawa and Iwata 2013a). Mainly based on molecu- lar evidence, the monophyly was only partially confirmed based on few species sequences of this group, and the as- semblage was confirmed to be part of the Acanthogobi- us-lineage (a monophyletic lineage of the family Gobiidae comprises about 29 genera) (Agorreta et al. 2013; Thacker and Dawn 2013). The phylogenetic position of S. fundico- Ja remained unknown up until now, for the sequence from S. fundicola was not included in these molecular studies. Since its original description, S. fundicola has occa- sionally been found in marine surveys (Kuroda 1957; Yamamuta et al. 1993; Shinohara et al. 2001; Shinohara et al. 2005; Iwatsuki et al. 2017; Choi and Lee 2019; So- noyama et al. 2020). So far, S. fundicola is mainly known from temperate regions of Japan, including off the Pacific coast from Matsushima Bay to Tosa Bay (Fig. 1, black arrows 2 and 3), Japan Sea coast from Aomori Prefec- ture to Yamaguchi Prefecture (Fig. 1, black arrows 4 and 5), and the Okinawa Trough (black full circle 6) (Akihito et al. 2013). This species is also reported off Tongyeo- ng, South Korea (Fig. 1 blue full circle 7) (Choi and Lee 2019). Although Oktyama (2014) recorded juveniles as 120°0'0"E 130°0'0"E 120°0'0"E 130°0'0"E S. fundicola from two stations of the East China Sea in 1956 (Fig. 1, orange full circle 8 and 9), this species iden- tification was not accepted by subsequent authors (Akihi- to et al 1984; Wu and Zhong 2008; Akihito et al 2013; Wu and Zhong 2021). So far, there is no record of this species from the Yellow Sea. According to the historical records, there are about 50 goby species known to occur in the Yellow Sea, all of which live in shallow coastal waters (Wu and Zhong 2008; Liu et al. 2015; Parenti 2021). Two R/V cruises, conducted during 2022, yielded four goby specimens from stations H27 (35°59.69'N, 123°07.63'E) and H12 (33°59.55'N, 123°22.99'E) in the Yellow Sea (see Fig. 1, green triangles). These specimens have notably large eyes, and combined with other mor- phometric characters these four gobies would conform to Suruga fundicola Jordan & Snyder, 1901. But there were a few inconsistent descriptive features, which hampered a definite identification. In order to confirm this morphol- ogy-based species identification, CO/ and /2S genes of mitochondrial DNA of these four specimens were ampli- fied. Phylogenetic analyses of the obtained sequences and the literature available sequences of other Acanthogobi- us-lineage gobies (also including one GenBank-retrieved 12S sequence of S. fundicola from Jogashima Island, Ja- pan, accession number LC069781), showed that the spec- imens belong to the species S. fundicola. This is the first report on the occurrence of S. fundicola in the Yellow Sea based on available specimens. 140°0'0"E = EAS} FU E 4v ; \ e 30°0'0"N 140°0'0"E Figure 1. Historical distributional records of S. fundicola and the sampling stations. The red full circle 1 indicates the type locality: Sagami Bay (Jordan and Snyder 1901); black arrows 2 and 3 indicate the distribution from Matsushima Bay to Tosa Bay, and black arrows 4 and 5 indicate the distribution from Aomori Prefecture to Yamaguchi Prefectural, black full circle 6 indicates Okinawa Trough (Akihito et al. 2013); blue full circle 7 indicates the records of Tongyeong of South Korea (Choi and Lee 2019); orange full circle 8 and 9 indicate the records in the East China Sea (Okiyama 2014); the green triangles H27 and H12 point to the locations of the survey stations where the four goby specimens were caught. zse.pensoft.net Zoosyst. Evol. 99 (2) 2023, 489-501 Materials and methods Specimen sampling and preservation The fisheries resources survey vessel Lanhai 101 carried out two R/V cruises conducted by the Yellow Sea Fisher- ies Research Institute (YSFRI) in spring (April) and sum- mer (July) 2022. A small Agassiz trawl (with a mouth of 1.8 m in width and 0.6 m in height) was employed during each cruise, at an average speed of 3 n mile/h (=5556 m/h) for 20 minutes. Four specimens of S. fundicola were col- lected, two in the spring (at H27 on 15 April) and two in the summer (one at H12 on 16 July and one at H12 on 20 July). The environmental parameters were obtained by the CTD measurements (SBE 911). Immediately after the capture of these specimens, the digital photographs were taken for each of them in a special glass tank with a Canon 5DSR, equipped with a micro lens (Canon), see Fig. 2. The muscle of the right lateral body was removed 491 and stored in 95% ethanol for further molecular analysis. The voucher specimens were then fixed in 10% forma- lin preservative and then transformed to 75% alcohol for morphological examination and permanent curation, and deposited in the National Marine Fishery Biological Ger- mplasm Resource Bank, China. Morphological analysis Meristic counts and morphometric measurements fol- lowed the methods used by Choi and Lee (2019). Addi- tionally, two other counts (Pelvic fin rays and Dorsal fin pterygiophore formula) and one morphometric measure- ment (Caudal peduncle length) were used. Thus, ten mer- istic and twenty-one morphometric measurements were taken for each individual in this study (Table 1). Measure- ments and counts were taken on the left side of the speci- mens whenever possible. Measurements were made point Table 1. Comparison of meristic and morphometric data of S. fundicola with the literature values. The numbers in bold point to the observed differences. Dorsal-fin pterygiophore formula is expressed as “3/I II II 11 I 0 1/12”, where “3” shows the number of vertebrae before the 1* dorsal fin is inserted, the Roman numerals in uppercase show the number of the dorsal pterygiophores inserted between the neural spines, the Roman numeral in lowercase shows the number of the interdorsal pterygiophores, and “12” shows that the two pterygiophore of the 1‘ ray of the second dorsal fin are mounted over the 12" vertebrae (Akihito et al. 1984; Shibukawa and Iwata 2013). Characteristic Present study Original description Choi and Lee (2019) Range Mean Standard length (mm) 51.8-63.5 58.7 Meristic counts First dorsal fin rays Vill Vill Second dorsal fin rays |, 16 |, 16 Anal fin rays |, 15-16 |, 15.8 Pectoral fin rays 20-21 20.3 Pelvic fin rays I-5 I-5 Lateral line scales 39-41 40 Transverse scales 8-10 9.0 Predorsal scales 10-11 10.8 Vertebral number 14+21 14+21 Dorsalfin pterygiophore formula 3/1 I110i/12 Morphometric data Percentage against SL (%) Head length 24.5-28.0 26.1 Head depth 13.1-17.6 15.4 Head width 13.2-14.2 13.6 Snout length 4.5-5.7 5:5 Eye diameter 8.1-9.7 8.7 Interorbital width 0.9-1.9 1.4 Jaw length 8.3-10.3 9.0 Body width 10.6-10.3 11.8 Body depth at origin of first dorsal fin 15.3-22.4 18.9 Body depth at origin of anal fin 16.0-18.3 16.9 Snout to origin of first dorsal fin 31.5-33.4 32D Snout to origin of second dorsal fin 53.3-58.7 55.1 Snout to origin of anal fin 55.7-61.1 59.1 Caudal peduncle length 10.7-13.5 11.7 Caudal peduncle depth 6.9-8.8 8.0 Pectoral fin length 19.5-21.8 20.6 Base of dorsal fin 13.8-14.7 14.3 Base of second dorsal fin 33.7-36.6 35.4 Base of anal fin 30.2-37.7 32.4 Caudal fin length 19.8 -23.3 21.4 Range Mean Range Mean 50.0-63.0 55.3 44.3-51.8 Vill Vill Vill Vill 17-19 17.8 |, 16-17 16-18 Ty |, 15-16 20-22 38-44 40.5 37-42 10-11 10-12 10.7 8-11 14+21-22 26.0-27.0 26:3 25.9-29.7 27.8 14.6-17.6 16.1 15.1-18.3 16.8 5.0-7.0 6.0 5.4-7.8 6.4 9.5-11.0 10.1 9.2-11.4 10.0 1.0 1.0 0.7-2.1 ies) 10.0-11.0 10.3 8.6-11.9 10.5 10.9-14.7 12.6 15.3-19.0 16.9 12.1-15.1 13.4 32.0-36.0 330) 28.6-36.0 33.4 52.0-54.0 O20 51.4-55.7 ete, 56.0-58.0 S75 53.0-58.6 pee 10.0-14.0 11.8 7.0-7.5 #1 6.7-9.1 8.2 21 21 17.7-20.9 19.0 13.4-16.3 14.7 30.8-38.3 35.6 30.6-37.0 32.4 18.6-27.2 22.3 zse.pensoft.net 492 Changting An et al.: Identification of the Suruga fundicola from the Yellow Sea to point with a digital caliper linked directly to a data-re- cording computer and the data were recorded to the near- est 0.1 mm. The measurements of the body were given as percentages (%) of the standard length (SL). Detailed in- formation about the meristic and morphometric measure- ments is provided in Table 1. Osteological features were observed with the help of micro-CT radiographs (Bruker, skyskan-1276) or from X-rays (Aolong, version 90). Ce- phalic sensory canals and papillae were recorded from specimens stained with cyanine blue following Akihito et al. (1984), the notations following Sanzo (1911); Shibu- kawa and Aizawa (2013); Shibukawa and Iwata (2013). DNA extraction, amplification and sequencing DNA was extracted by using TIANamp Genomic DNA Kit (Tiangen Biotech, Being) according to the man- ufacturer’s recommended protocol, and the quality was estimated at wave-length 260/280 nm by a Nano- 300 micro-spectrophotometer (Allsheng, Hangzhou, China). The obtained DNA solutions were stored at -20 °C until used. The mitochondrial /2S rRNA (/2S) and the cyto- chrome c oxidase I (COI) genes were amplified by PCR with different primer combinations. For /2S, we used MiFish-U-F and MiFish-U-R designed by Miya et al. (2015). Primers reported by Ward et al. (2005) were used for the amplification of CO/. PCR was conducted in 25 ul volumes, including 12.5 ul Master mix Taq, 1 ul of each primer, 1 pl template DNA, adding double distilled water to adjust the volume. Thermocycling con- ditions were as follows: initial denaturation for 4 min at 94 °C, denaturation for 50 s at 94 °C, annealing for 50 s at 55 °C for CO/ and 59 °C for /2S, and extension 1 min for COI and 30 s for /2S at 72 °C. After 35 cy- cles, the final extension was done at 72 °C for 10 min. The PCR products were bidirectional sequenced by BGI Genomics Co., Ltd.. In this study, four specimens of S. fundicola and three specimens of Amblychaeturichthys hexanema (Bleeker, 1853) were sequenced for phyloge- netic analysis, and all amplified CO/ and /2S sequences were submitted to GenBank (for the accession numbers, see Table 2). Molecular sequence analysis All amplified sequences of the two mtDNA genes were concatenated and used for molecular phylogenetic analy- sis, along with 37 GenBank-retrieved sequences from 19 related species of 14 genera belonging to the Acanthogo- bius-lineage. In addition, Odontobutis haifengensis Chen, 1985 (Odontobutidae) was used as an outgroup (Table 3). Multiple alignments were prepared for CO/ and /2S se- quences using the program MUSCLE itn MEGA X (Edgar Table 2. The detailed information of specimens analyzed in this study. Species Specimen catalog GenBank no. Sampling location Resource col 12S rRNA Suruga fundicola 1 YSFRI27216 OP824753 OP837791 Yellow Sea station H27 Present study S. fundicola 2 YSFRI27217 OP824754 0OP837792 Yellow Sea station H27 Present study S. fundicola 3 YSFRI36942 OP824752 OP837789 Yellow Sea station H12 Present study S. fundicola 4 YSFRI36943 OP824755 OP837790 Yellow Sea station H27 Present study S. fundicola 5 CBM:ZF:15688 ve LC069781 Japan: off west of Jogashima Island GenBank Amblychaeturichthys YSFR27208 OP824756 OP837786 North Yellow Sea Present study hexanema 1 Am. hexanema 2 YSFR27209 OP824757 OP837787 North Yellow Sea Present study Am. hexanema 3 YSFR27210 OP824758 OP837788 North Yellow Sea Present study Am. hexanema 4 Uncatalogued KT781104 KT781104 China: Qingdao, Shandong Prov. GenBank Acanthogobius flavimanus Uncatalogued MW271007 MW271007 Uncatalogued (Maybe from China) GenBank Ac. hasta Uncatalogued MK253669 MK253669 China: Lianyungang City, Jiangsu Prov. GenBank Chaeturichthys stigmatias Uncatalogued MNO38166 MNO38166 China: Qingdao, Shandong Prov. GenBank Lophiogobius ocellicauda Uncatalogued KR815520 KR815520 China: Zhoushan, Zhejiang Prov. GenBank Lepidogobius lepidus UW:151092 KF918879 LC092050 USA: Washington, Puget Sound GenBank Chaenogobius gulosus JM120726-11 KP696748 KP696748 Korea: coastal area of Jangmok GenBank Chaenogobius annularis Uncatalogued OM830225 OM830225 China GenBank Gymnogobius urotaenia Uncatalogued KT601093 KT601093 Uncatalogued (Maybe from South Korea) GenBank Parachaeturichthys polynema ECSFRIENMWO1 OKO12405 OK012405 China: East China Sea GenBank Eucyclogobius newberryi LodgeLab Enewberryi_l KPOQ13101 KPO13101 Uncatalogued GenBank Gillichthys mirabilis Uncatalogued FJ211845 FJ211845 China: Nantong city, Jiangsu Prov. GenBank Gymnogobius petschiliensis 20131115NA05 AY525784 AY525784 China: Qingdao, Shandong Prov. GenBank Luciogobius platycephalus Uncatalogued JX971538 JX971538 China: Zhoushan, Jiangsu Prov. GenBank L. pallidus Uncatalogued KF040451 KFO40451 South Korea: Jeju Island GenBank Tridentiger bifasciatus Uncatalogued JN244650 JN244650 China: Zhoushan fishing ground Zhejiang Prov. GenBank T. trigonocephalus Uncatalogued KIZ262115-. *KI282115 Uncatalogued (Maybe from China) GenBank Rhinogobius similis Uncatalogued KF371534 KF371534 — China: Liangzi Lake in the middle reaches of the GenBank Yangtze River Odontobutis haifengensis Uncatalogued MF383619 MF383619 China: Fengshun, Guanggong GenBank (Odontobutidae) zse.pensoft.net Zoosyst. Evol. 99 (2) 2023, 489-501 Table 3. The depth (D), temperature (T), and salinity (S) of H12 and H27. Data Station Bottom layer Surface layer D(m) TCC) S(%o) D(m) TCC) SM%o) Aphlrl6<2022." “HI2* =69;0" 105° 33,0: siz AZ: 132.6 April’t5,2022* .H27 (74:0. 9.5 “328 2:8 10:5. 332:6 July 16, 2022 Has, 69:0: TWOs6. 53: 2:0: “269: 308 July 20, 2022 2A ey PAO! ce Oe2t oe3228 2B 25,6-— 30.8 2004). Genes were concatenated with the help of Sequen- ceMatrix 1.8 (Vaidya et al. 2011). The genetic distance was calculated in MEGA X, based on Kimura 2-parame- ter (K2P) (Kimura 1980), and the base composition was calculated with the same software, based on K2P (Sudhir et al. 2018). The optimal evolution model was GTR se- lected in MEGA X based on Akaike’s information crite- rion (AIC), and the maximum likelihood tree (ML tree) was constructed with the same software (Kumar et al. 2018), with 1,000 bootstrap replications. Results Suruga fundicola Jordan & Snyder, 1901 Figs 2, 3, Table 1 Suruga fundicola Jordan & Snyder, 1901: 96, fig. 20 (original descrip- tion, type locality: Sagami Sea, Japan); Akihito et al. 1984: 279 (in English), fig. 253-H; Akihito et al. 2002: 1207 (in Japanese); Akihito et al. 2013: 58 (in Japanese); Shibukawa and Iwata 2013a: 45; Mat- sul et al. 2014: 6; Choi and Lee 2019:255, fig. 1. Diagnosis. Distinct from all other gobies (Gobiidae), members of the Acanthogobius-group share a unique dominant pattern of the dorsal-pterygiophore formula, 3/I 1 WIT I 0 (Akihito et al. 1984). In the Acanthog- obius-group, S. fundicola can be distinguished from the species of Sagamia, Siphonogobius and Pterogobius by possessing no free rays in the upper part of the pectoral fin and the posterior margin of the pelvic frenum indent- ed. S. fundicola can be distinguished from the species of Lophiogobius, Amblychaeturichthys, and Chaeturichthys by the lack of barbels or flaps on the ventral surface of the head (except for the mental frenum). From species of Acanthogobius, S. fundicola can be distinguished by the large eye, its diameter greater than the snout length (vs. usually less); each cephalic sensory papilla formed into a minute skin flap (vs. not), the posterior oculoscapular canal absent (vs. posterior oculoscapular canal and its ter- minal pores K’ and L’ present). Description of Yellow Sea specimens. The counts and measurements are given in Table 1. Dorsal-fin rays VIII-I, 16; anal-fin rays I, 15 (1), I, 16 (3); pectoral-fin rays 20 (3), 21 (1); pelvic fin rays I, 5 (4); longitudinal scales 39 (1), 40 (2), 41 (1); pre-dorsal mid-line scales 10 (1), 11 (3); transverse scales 8 (1), 9 (2); 10 (1); vertebral count 14+21 = 35 (4); dorsal-pterygiophore formula 3/I II 493 IT 1110 1/12; epural 2; anal-fin pterygiophores anterior to first haemal spine 2. The following measurements are in % SL: head length 24.5—28.0 (mean 26.1); head depth 13.1-17.6 (15.4); head width 13.2—14.2 (13.6); snout length 4.5—5.7 (5.5), eye diameter 8.1—9.7 (8.7); interorbital width 0.9-1.9 (1.4); jaw length 8.3-10.3 (9.0); body width 10.6-10.3 (11.8); body depth at origin of first dorsal fin 15.3—22.4 (18.9); body depth at origin of anal fin 16.0-18.3 (16.9); snout to origin of first dorsal fin 31.5 —33.4 (32.5); snout to origin of second dorsal fin 53.3—58.7 (55.1); snout to origin of anal fin 55.7-61.1 (59.1); caudal peduncle length 10.7—13.5 (11.7); caudal peduncle depth 6.9-8.8 (8.0); pectoral fin length 19.5—21.8 (20.6); base of dorsal fin 13.8-14.7 (14.3); base of second dorsal fin 33.7—36.6 (35.4); base of anal fin 30.2—37.7 (32.4); caudal fin length 19.8—23.3 (21.4). General body appearance was shown in Figs 2, 3. Body small, moderately elongated; predorsal body profile slightly convex; ventral profile slightly concave, espe- cially from pectoral-fin insertion to anal-fin origin. Head large, not depressed, short but longer than wide, depth and width less than those of the body. Snout short, obtuse in lateral and dorsal view, shorter than eye diameter and postorbital head length. Eyes notably large, situated dor- solateral in upper half of the head, with very narrow in- terorbital space, eyes nearly meeting, diameter larger than interorbital space or snout length. Mouth almost terminal, but upper jaw slightly protruding. Maxillary concealed except at its posterior end. Tongue thick, rather broad, round anteriorly. Gill openings broad, extending anteri- orly to the vertical line of the posterior margin of the eye; upper edge of the gill opening on fleshy pectoral-fin base, slightly above the upper margin. No barbels. Body cov- ered with cycloid scales, anterior small, posterior large and the scales are rather loosely attached. Head naked. Fins flexible, without spinous rays. First dorsal fin with 8 slender spines, reaching origin of second dorsal when depressed; dorsal-pterygiophore formula 3/I I III 1 0 1/12. Second dorsal fin with 1 simple and 16 branched rays, shorter than the first spines. Origin of first dorsal fin posterior to a vertical through base of pectoral fins, first dorsal fin without filamentous spines. The distal margin of the first dorsal fin is convex, when adpressed, the dis- tal tip touches the base of the spine of the second dorsal fin. Dorsal fins discontinuous. Origin of second dorsal fin somewhat at vertical through the anus, and anterior to the anal fin. When adpressed, the distal tips of the second dorsal fin and the anal fins do not reach the procurrent rays of the caudal fin. Pectoral fins rounded, with 20 rays. The pectoral fin extends posteriorly to the vertical line through the posterior margin of the base of the first dorsal fin. Pelvic fin fused into a disc, each with 1 simple and 5 branched rays. Anal fin with 15-16 rays, the anterior of the anal fin below the third branched dorsal ray of the second dorsal fin. Segmented caudal-fin rays 7+7, upper unsegmented caudal fin rays about 12 and lower unseg- mented caudal fin rays about 11. zse.pensoft.net 494 Changting An et al.: Identification of the Suruga fundicola from the Yellow Sea Figure 3. Lateral (a), dorsal (b), and ventral (¢c) views of S. fundicola: YSFRI27216, 63.5 mm SL. Cephalic canals are variably developed and are shown in Fig. 4: anterior oculoscapular canal (AOC) with B', D (S), FE, H’; posterior oculoscapular canal (POC) absent; preopercular canal (PC) with pores M' and O’; four short longitudinal sensory papillae (SSP) rows (=rows r, u, s, t) on snout; four SSP rows (=rows g, j, k, and 1) close behind zse.pensoft.net the eye; two SSP rows (=rows h, 1) before dorsal fin; two transverse sensory papillae (TSP) rows (=rows n and o) on snout and behind the eye, respectively; four longitu- dinal sensory papillae (LSP) rows (=rows a, b, c, and d) on the cheek; anterior end of row a approaches the ante- rior margin of the eye; rows b and c very close together; Zoosyst. Evol. 99 (2) 2023, 489-501 Figure 4. Dorsal (top), lateral (middle), and ventral (bottom) views of the head of S. fundicola. YSFRI27216, 63.5 mm SL female), showing cephalic sensory canal pores (indicated by roman uppercase letters, except for AN and PN) and papillae (indicated by roman lowercase letters). AN and PN, indicated anterior and posterior nares, respectively. row cp with a single sensory papilla; row d arc-shaped, extending posteriorly to the vertical line through the pos- terior margin of the pupil; two long parallel longitudinal rows of sensory papillae just behind the chin (=row f), and ending on both sides at the opercles, one TSP row (=row ot) and two LSP rows (=rows os and 01) on the opercles, row ot extends to the ventral side. Cranium flat, frontals extremely narrow (Fig. 5a, e). No suborbital bone. Five branchiostegal rays, the first one thin, and last one strong (Fig. 5f). Four pairs of cer- atobranchials (Fig. 5g). Well-developed teeth on upper and lower pharyngeal. Three pairs of otoliths, sagittae, lapillus and asteriscus (Fig. 5e). Vertebral count 35, 14 abdominal vertebrae (av) and 21 caudal vertebrae (cv), 14 pairs of ribs appending on parapophysis (Fig. 5a, b). Three hypurals (HY), respectively HY1+2 (HY1 and HY2 fused into one), HY3+4 (HY3 and HY4 fused into one), and HY5; two epurals, EP] and EP2. Coloration. In freshly collected specimens (Fig. 2), head and dorsum of body dusky, darker on snout, with several irregular light-yellow blotches on the lateral body, 495 ventral body lighter, abdomen almost white. Pupil of the eye black, iris golden gray. A light sapphirine blotch pres- ent on the gill cover. Six or seven large dark spots scat- tered along middle of the side from the gill opening to the caudal-fin base; 2 or 3 light orange stripes on gray dorsal and caudal fins, the anterior margin of first dorsal fin with dusky spots, the upper posterior of caudal fin with a black stripe, anal fin somewhat gray. Coloration changed after 2 months of preservation (10% formalin preservative and then transformed to 75% alcohol), the yellow and orange pigment disappeared from body and fins, and the body of the fish became dark-yellowish, covered with tiny black spots, back dark- er and belly lighter, snout black, lateral dark spots not clear. Pupil of the eye white, iris golden black. Dorsal, pectoral, pelvic and anal fins light greyish. Distribution. Northwest Pacific: off Pacific coasts from Miyagi Prefecture to Tosa Bay, Japan Sea from Aomori to Yamaguchi Prefecture, Okinawa Trough (Akihito et al. 2013), Southern Sea of Korea (Choi and Lee 2019), East China Sea (Oktyama 2014) and Yellow Sea (present study). Habitat and ecology. The four specimens were col- lected at depths between 69 and 74 meters (Fig. 1). The two stations maintained a relatively low temperature of about 10 °C and a high salinity of about 33%o in April and July 2022 (Table 3). This species is considered as one of the deepest dwelling goby in Japan, known from depths of 40 to 400 meters (Akihito et al. 2013; Choi and Lee 2019). The catch at the stations mainly consists of ophiuroids, molluscs, jellyfishes, fishes and so on, most common spe- cies of which are the brittle stars Ophiura sarsii vadicola Djakonov, 1954 (Ophiuroidea) and Stegophiura sladeni (Duncan, 1879) (Ophiuroidea) (Fig. 6). Examples of the co-existing fish species are Jaydia lineata (Temminck & Schlegel, 1843) (Apogonidae), Cleisthenes pinetorum Jordan & Starks, 1904 (Pleuronectidae), Liparis tanakae (Gilbert & Burke, 1912) (Liparidae), Pholis fangi (Wang & Wang, 1935) (Pholidae), and Hexagrammos otakii Jor- dan & Starks, 1895 (Hexagrammidae). Sequence characteristics and phylogenetic place- ment. The concatenated CO/ and /2S sequences from 22 species were 704 bp in length (after trimming, except LC069781), including 400 conserved sites, 307 variable sites, 278 parsimony informative sites, and 24 singleton sites. The mean four nucleotide frequency of S. fundicola was A=26.1%, T=28.8%, C=27.3% and G=17.8%, slight- ly A-T rich (54.9%). The intragroup sequence divergence of S. fundicola was 0.5%; the genetic distance between samples of the Yellow Sea and the sequence (LC069781) of S. fundicola from west of Jogashima Island of Japan was 0.2%. This species has a genetic distance of 19.2% (C. stigmatias) to 26.3% (E. newberryi) to the other 20 Species we used (see Table 4). The ML tree based on the concatenated sequences 1s shown in Fig. 7. In the tree to- pology, all species from the same genus clustered in one lineage; the four sequences of S. fundicola clustered into a highly supported (94% bootstrap P value) lineage and zse.pensoft.net 496 Changting An et al.: Identification of the Suruga fundicola from the Yellow Sea _ J a i _ ~~ a . ms tae 5 ag a i] ; ~~ ’ ‘ , aX) : Ltt MT AM ~ << - - —, —_— a 28 29 30 31 32 Figure 5. Micro-CT images of right (a), dorsal (b), and ventral (c) views of specimen YSFRI27216; front (d) dorsal (e), ventral (f), and oblique view (g) of the head of specimen YSFRI27216. 1. teeth, 2. premaxilla, 3. maxilla, 4. palatine, 5. ectethmoid, 6. paras- phenoid, 7. frontal, 8. parietal, 9. supraoccipital, 10. neural spine, 11. first dorsal fin spines, 12. interdorsal pterygiophores, 13. pte- rygiophore, 14. second dorsal fin rays, 15. neural spine of preural centrum 3(NPU3), 16. neural spine of preural centrum 2(NPU2), 17. epural 1 (EP1), 18. epural 2 (EP2), 19. urostyle, 20. hypural 5 (HY5), 21. hypural 3+4 (HY3+4), 22. hypural 1+2 (HY1+2), 23. dental, 24. articular, 25. sympletic, 26. branchiostegal rays, 27. preopercular, 28. subopercular, 29. proximal radials, 30. pectoral fin soft rays, 31. pelvic fin spine, 32. vertebral canal, 33. boundary of abdominal vertebra and candal vertebrae, 34. anal fin rays, 35. ventrispinales, 36. haemal spine of preural centrum 3 (HPU 3), 37. haemal spine of preural centrum 2 (HPU 2), 38. parhypural (PH), 39. caudal fin ray, 40. rib, 41. parapophysis, 42. pelvic bone, 43. ethmoid, 44. lapillus 3+4 (HY3+4), 45. sagittae, 46. asteriscus, 47. basihyal, 48. ceratohyal, 3 (HPU 3). 49. epihyal, 50. cleithrum, 51. pharyngeal tooth, 52. ceratobranchial. zse.pensoft.net Zoosyst. Evol. 99 (2) 2023, 489-501 cae ee i Figure 6. The catch of A: H27 (15 April), B: H12 (15 July), and C: H27 (20 July). 100 KP696748 Chaenogobius gulosus OM830225 Chaenogobius annularis JX971538 Luciogobiusplatycephalus KF040451 Luciogobius pallidus AY525784 Gymnogobius petschiliensis 100 KT601093 Gymnogobius urotaenia UW:151092 Eucyclogobius newberryi KF918879 Lepidogobius lepidus FJ211845 Gillichthys mirabilis JN244650 Tridentiger bifasciatus KT282115 Tridentiger trigonocephalus KF371534 Rhinogobius similis OKO012405 Parachaeturichthys polynema KC480264 Lophiogobius ocellicauda MW271007 Acanthogobius flavimanus MK253669 Acanthogobius hasta 30 ) YSFRI36942 Suruga fundicola* YSFRI27216 Suruga fundicola* YSFRI36943 Suruga fundicola* YSFRI27217 Suruga fundicola* MN038166 Chaeturichthys stigmatias YSFRI27209 Amblychaeturichthys hexanema* 100 KT781104 Amblychaeturichthys hexanema 84 YSFRI27210 Amblychaeturichthys hexanema* 99 | YSFRI27208 Amblychaeturichthys hexanema* MF383619 Odontobutis haifengensis 70 68 46 69 96 96 18 100 61 55 47 100 87 100 37 | LC069781 Suruga fundicola 94 99 rH 0.050 Figure 7. ML tree inferred from concatenated CO/ and /2S sequences. Numbers at major internal nodes are bootstrap probability values. Sequences with * were sequenced in the present study. had a sister group relationship with the lineage formed by A. hexanema and C. stigmatias. Material examined. YSFRI27216—27217, 2 spec- imens, 51.2-63.5 mm SL, station H27, Yellow Sea, off Qingdao, Shandong Province, China (35°59.69'N, 123°07.63'E), collected by Changting An on 15 April, 2022; YSFRI36942, 1 specimen, 60.5 mm SL, station H12, Yellow Sea, off Lianyungang, Jiangsu Province, China (33°59.88'N, 123°24.14'E), collected by Hongyue Sun, on 16 July, 2022; YSFRI36943, 1 specimen, 59.1 mm SL, station H27, Yellow Sea, off Qingdao, Shandong Province, China (35°56.03'N 123°07.54'E), collected by Hongyue Sun, on 20 July, 2022. Discussion According to Jordan & Snyder’s (1901) record, the type specimen (USNM 49744) was caught in a depth of 65 fathoms (119 meters), off Sagama, Japan. Unfortunately, the holotype of this species cannot be examined now, for it was lost in 1980 (Fricke et al. 2023). Our photographic zse.pensoft.net 498 Changting An et al.: Identification of the Suruga fundicola from the Yellow Sea Table 4. Genetic distances (%) based on concatenated CO/J and /2S sequences computed by MEGA X among 21 groups. S. fundi- cola * was sequenced for the present study. Group Intragroup Intergroup | ie meees Sage cts eg rc ameents meal Ge Nid ued ARS Reel tesaciBs md hoy mally = AA pieN bc? et10) 1. Suruga fundicola * 0.5 2. S&. fundicola n/c 0.2 3. Am. hexanema 0.5 Alr5* 1955 4 Ac. flavimanus n/c 23.9 26.5 26.6 5 Ac. hasta n/c 24.4 22.9 23.7 13.6 6 C. stigmatias n/c 192 TASe lS. 26.6 22:8 7 Lo. ocellicauda n/c 216° 201° 273° 21e2. 20.9°75.5 8 Ch. gulosus n/c 23.5 14.8 26.6 24.6 25.2 23.4 20.2 9 Ch. annularis n/c 23.9 16.1 25.7 26.3 25.3 25.0 25.3 11.2 10 Gy. urotaenia n/c 21,4. 18:7 25.0° 24.3: 22:6 23:77 23.6. 16:2 15:4 11 Gy. petschiliensis n/c 23:5. 22:2025.9 26:0.23.8625.3 -2410.16.9- 116,29 2:8 12 P. polynema n/c 24.0 21.8 27.9. 22:9. 26.1. 26.3°21.3-27.0'28)5-23:9 23.1 13 E. newberryi n/c 26.3: 22.6 31.4 25.0 24.5 28:2 26.8 19.7 21.6 21.0 21.8: 26.9 14. Gi. mirabilis n/c 20.9 12.6 23.9 24.7 21.4 23.3 20.6 17.4 21.4 17.1 18.7 23.4 20.1 15 Le. lepidus n/c 21.1 16.1 24.9 24.1 22.9 22.0 24.3 19.2 20.2 16.0 16:9 24.7 17.2 16.2 16 Lu. platycephalus n/c 21.4 15:5 24.5 23.2 23.8°23.6 22:2 13.7 14.0 14.1. 14.8°23.3 19.3 15.6 17.5 17 Lu. pallidus n/c 22.8; 20? -2?.8-23,5°22.3: 21.2 24,7 154A 15.4 13.1 14.0:24.6 19.3 15:9. 16:5° 10:9 18 T. bifasciatus n/c 25.0 29.0 26.2 26.3: 25.4 -24.7 24.5.24.4 25.1 23.2 23.5°25.0°24.4 24.7 25.3. 25:1 24.6 19 T. trigonocephalus n/c 23.7 29.0 26.2 25.4 25.8 26.9 26.8 27.5 26.1 24.8 24.8 24.3 28.0 24.6 25.5 24.0 24.8 12.9 20 R. similis n/c 20.1 14.8 23.0 23.8 24.0 22.2 23.3°19.4.21.4: 18.7 19.7 22.2 22.9 18:6 19,2 18.3 °20.0 21.1 21.9 21 O. haifengensis n/c 22:5 23:5-27.1 24.1 26.2 26:3°24.5 25.7°26.4-22.4 21.8.24.1 24.8 24.4 22.8 22.7 20:9 29,7 27.3 22.3 examination based on two paratypes AMNH 1-3549 (48.2-60.3 mm SL, Fig. 8) and their X-ray images, found that this species has notably large eyes, a very narrow interorbital space, jaws equal, eight dorsal spines and 17— 18 dorsal-fin branched rays, dorsal-pterygiophore formu- la 3/1 If UW 11101/12, 17-18 anal-fin branched rays, and 35 (14 AV and 21 CV) vertebrae, as stated in its original description. All these characters are shared by the four specimens collected from the Yellow Sea of China, which are therefore conspecific with this species. Clearly, the four specimens AMNH I-35829 (91.40- 104.55 mm SL, Fig. 9), caught by Pope from Foochow (=Fuzhou), Fukien (=Fuyian), China, were misidentified as they have relatively small eyes (less than the snout length), a broad interorbital space, a lower jaw projecting beyond the upper jaw, eight spines of the first dorsal fin and 16-17 branched rays of the second dorsal fin, the dorsal-fin pte- rygiophore formula 3/1 II IIT 110 1/11, 17-18 anal fin rays, and 34 (13 AV and 21 CV) vertebrae. Based on the num- ber of rays in the second dorsal finI-16 (4), they can be distinguished from species of Chaeturichthys (1-20-22), Siphonogobius (1-12—13), Pterogobius (1-18-27), and Ac- anthogobius (1-18-20). Based on the number of anal fin rays (17-18), they are distinct from species of Sagamia (13-14), and Amblychaeturichthys (11-13). In compari- son to S. fundicola, the eye diameter is clearly less than the snout length (vs. longer). To sum up, all the above-men- tioned characters of the four specimens AMNH I-35829 are consistent with Lophiogobius ocellicauda Gunther, 1873, as recorded by Wu and Zhong (2008). Despite the recognition of four specimens from the Yellow Sea as S. fundicola, the following characteristics slightly differ from the data given in the original descrip- tion. The head depth and width were less than those of the body (vs. head deeper and broader than those of the body). zse.pensoft.net Based on Fig. 2, the dorsal fin has two or three light orange stripes (vs. fins dusky) and the head and dorsum of the body are dusky with several yellow markings (vs. dusky above). This body coloration is similar to that described by Shibukawa and Aonuma (2007). Nonetheless, the sam- pling location of the sequence (LC069781) is Kanagawa, Muura, off the west of Jogashima Island, very close to the type locality of S. fundicola given a 0.5% sequence diver- gence between this sequence and four sequences from the Yellow Sea samples, the voucher specimens of the four sequences can be recognized as the species S. fundicola. The aforementioned variations are probably intraspecific. A total of 15 goby specimens from Tongyeong of South Korea were recognized as S. fundicola by Choi and Lee (2019). Most characteristics of the specimens are consis- tent with our examination of the specimens from the Yellow Sea. This study disagrees with Choi and Lee’s (2019) report in two characters: head width and body depth at the anal-fin origin (Table 1). These variations can be caused by many factors, such as size-related, specimen condition, measur- ing method, and so on, which needs further examination. It is worth mentioning that our samples from the Yellow Sea of China (51.8-63.5 mm SL) are a bit larger than Choi and Lee’s (2019) specimens of this species (44.3—51.8 mm SL). The larval specimens, collected from two stations of the East China Sea by beam trawl in June 1956 [orange full circle 8 (32°04'N, 123°03'E) and 9 (31°53'N, 123°26'E); Fig. 1], were identified as S. fundicola (Oktyama 2014). We have no access to these juvenile specimens, so the possibility cannot be ruled out that they were misidenti- fied. But, considering that the sampling locations are not far from station H12, we believe that this historical record is credible. The species is regarded as the deepest dwelling Go- biidae in Japan, at depths from 40 to 400 meters with Zoosyst. Evol. 99 (2) 2023, 489-501 RoNbtiantg a" aaa) es Ly gl Ww 4 | c \ a ‘ 7 ‘VMSA be Oy: (Ns a lL Ne: ~~ — ~ > J Figure 8. Digital photos (a) and X-ray images (b) of the lateral body in paratypes of S. fundicola AMNH 3549, 48.2—60.3 mm SL, collected by D.S Honshu from Island, Japan. Figure 9. Digital photos (a) and X-ray images (b) of the lateral body in four goby specimens AMNH 35829, 91.40—104.55 mm SL, collected by Pope from Fuzhou, Fujian Province. zse.pensoft.net 500 Changting An et al.: Identification of the Suruga fundicola from the Yellow Sea a sandy silt bottom (Akihito et al. 2013; Choi and Lee 2019). The species was only discovered in two stations (H12 and H27, with a depth of about 70 m) of this sur- vey, with a relatively low temperature of about 10 °C in summer. Moreover, many studies indicate that the Yellow Sea Cold Water Mass (YSCWM) can help the low-tem- perature species to escape the high temperature stress in the summer, by providing an appropriate over-summer- ing habitat (Zhu et al. 2018; Li et al. 2021). However, up until now, there is no study indicating that S. fundicola is a true low-temperature species. Considering the water depth and temperature of its habitat, 1t is inferred that it must have some biological adaption to help it to live in this habitat. This goby species can possibly be used as an ideal experimental model for the study of adaptive evolu- tion in a population of deep water gobies. Suruga fundicola was assigned to the Acanthogobi- us-group based on morphological evidence (Birdsong et al. 1988; Shibukawa and Iwata 2013). But there was no molecular phylogenetic study which included a sequence of S. fundicola. This study also represents the first effort to address the issue about the relationship of Suruga with- in the Acanthogobius-lineage. The phylogenetic trees, based on concatenated CO/ and /2S genes, indicate that S. fundicola has a close relationship with A. hexanema, and C. stigmatias. Based on genetic distances of the concate- nated genes in our admittedly limited sampling, S. fundi- cola has the smallest genetic distances with C. stigma- tias (19.2%). In the present study, 6 species from the Acanthogobius-group were clustered into two different lineages, opposite of the result of Shibukawa and Iwata (2013a). Maybe the nucleotide sequences used by us are too short and somewhat uninformative to provide a suf- ficient phylogenetic signal. Therefore, it is necessary to conduct more phylogenetic studies with the help of data sets based on combined nuclear and mitochondrial genes from more species to provide a more realistic insight into the phylogeny of the Acanthogobius-lineage (Gobiidae). Comparative material Suruga fundicola. AMNH 1-3549, paratypes, 2 speci- mens, 48.2—60.3 mm SL; 1900; Suruga bay, Honshu Island, Japan (photograph examined). Lophiogobius ocellicauda. AMNH_ 1-35829, 4 speci- mens, 91.40-104.5 mm SL; Mar 1926; Foochow (=Fuzhou), Fukien (=Fujian Province), China (pho- tograph examined). Amblychaeturichthys hexanema: YSFRI27207—27210, 4 specimens, 60.9-85.5 mm SL; Sept 2022; Qiangdao, Shandong Province, China. Chaeturichthys stigmatias: YSFRI 34428-34429; 12 spec- imens; 68.9-95.5 mm SL; Dandong, Liaoning Prov- ince, China. YSFRI 34407-34408; 2 specimens; 68.9— 70.8 mm SL; Dandong, Liaoning Province, China. Acanthogobius hasta: YSFRI 34422-34431; 10 specimens; 67.5—-98.5 mm SL; Dandong, Liaoning Province, China. zse.pensoft.net Acknowledgments Our sincere thanks should be given to Mingwei Zhang (Ocean University of China, Qingdao, China), who shared the specimens with us, and the whole staff of the Lanhai 101 for their help. Especially grateful to Prof. E Zhang of Insti- tute of Hydrobiology, Chinese Academy of Sciences, Wu- han, China (THB), who provided constructive suggestions for this manuscript, and Dongming Guo for taking X-radio- graphs and Micro-CT images. Thanks to Xiao-dong Bian (Yellow Sea Fisheries Research Institute, Chinese Acade- my of Fishery Sciences), for providing valuable advice and valuable materials. Thanks should be given to Junsheng Zhong, who provided kindhearted guidance for the obser- vation of sensory canals and papillae. We thank Radford Ar- rindell (AMNH) for friendly help in providing photographs and X-radiographs. Special thanks go to Xiao Chen (Anhui Agricultural University), who provided friendly help with this manuscript. This research was funded by the National Key R&D Program (2018 YFD0900803), National Marine Aquatic Germplasm Bank Project, Central Public-interest Scientific Institution Basal Research Fund, YSFRI, CAFS (NO.20603022022024). Data and samples were collected onboard of R/V “Lanhai 101” implementing the open re- search cruise NORC2022-01 supported by NSFC Shiptime Sharing Project (project number: 42149901). Shufang Liu and Zhimeng Zhuang contributed to the design of the study. Shufang Liu supervised, reviewed, and edited the manuscript. Hongyue Sun, Changting An, and Kaiying Liu participated in the collection of speci- mens. Ang Li, Huang Wang and Busu Li provided con- structive suggestions for this manuscript. Changting An analyzed the data and drafted the manuscript. Richard van der Laan provided many scientific suggestions and improved the English writing. All authors contributed to the writing of the paper. All procedures described in this paper were in accor- dance with Chinese laws and were licensed by the Minis- try of Ecology and Environment of the People’s Republic of China. 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